International Journal of Aquaculture 2024, Vol.14, No.4 http://www.aquapublisher.com/index.php/ija © 2024 AquaPublisher, registered at the publishing platform that is operated by Sophia Publishing Group, founded in British Columbia of Canada. All Rights Reserved. Publisher
International Journal of Aquaculture 2024, Vol.14, No.4 http://www.aquapublisher.com/index.php/ija © 2024 AquaPublisher, registered at the publishing platform that is operated by Sophia Publishing Group, founded in British Columbia of Canada. All Rights Reserved. Publisher Aqua Publisher Editedby Editorial Team of International Journal of Aquaculture Email: edit@ija.aquapublisher.com Website: http://www.aquapublisher.com/index.php/ija Address: 11388 Stevenston Hwy, PO Box 96016, Richmond, V7A 5J5, British Columbia Canada International Journal of Aquaculture (ISSN 1927-5773) is an open access, peer reviewed journal published online by AquaPublisher. The journal publishes all the latest and outstanding research articles, letters and reviews in all working and studying within varied areas of aquaculture, containing the latest developments and techniques for practice in aquaculture; information about the entire area of applied aquaculture, including breeding and genetics, physiology, aquaculture-environment, hatchery design and management, utilization of primary and secondary resources in aquaculture, production and harvest, the biology and culture of aquaculturally important and emerging species. All the articles published in International Journal of Aquaculture are Open Access, and are distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. AquaPublisher uses CrossCheck service to identify academic plagiarism through the world’s leading plagiarism prevention tool, iParadigms, and to protect the original authors' copyrights. Aqua Publisher is an international Open Access publisher specializing in the field of marine science and aquaculture registered at the publishing platform that is operated by Sophia Publishing Group (SPG), founded in British Columbia of Canada.
International Journal of Aquaculture (online), 2024, Vol. 14 ISSN 1927-6648 http://aquapublisher.com/index.php/ija © 2024 AquaPublisher, registered at the publishing platform that is operated by Sophia Publishing Group, founded in British Columbia of Canada. All Rights Reserved. Latest Content 2024, Vol. 14, No.4 【Research Perspective】 Ecological Impacts of Common Carp Invasions: A Global Perspective 174-183 Qiong Wang, Chenmin Sun, Liping Liu DOI: 10.5376/ija.2024.14.0018 【Review Article】 Molecular Mechanisms Underlying the Diversification of Aquatic Life Forms 184-194 ManmanLi DOI: 10.5376/ija.2024.14.00019 【Research Insight】 The Role of Digital Technologies in Supporting Climate Change Adaptation in Fisheries and Aquaculture 195-210 Durdarshi Juggoo, Pierre Olivier St Flour DOI: 10.5376/ija.2024.14.0020 【Feature Review】 Sustainable Cultivation and Environmental Impact of Laminaria japonicaFarming 211-220 Peiming Xu, Xianming Li DOI: 10.5376/ija.2024.14.0021 Climate Change and Aquatic Ecosystem Health: Impacts, Adaptation Strategies, and Future Challenges 221-231 Xuelian Jiang, Wenfang Wang DOI: 10.5376/ija.2024.14.0022
International Journal of Aquaculture, 2024, Vol.14, No.4, 174-183 http://www.aquapublisher.com/index.php/ija 174 Research Perspective Open Access Ecological Impacts of Common Carp Invasions: A Global Perspective Qiong Wang, Chenmin Sun, Liping Liu Tropical Marine Fisheries Research Center, Hainan Institute of Tropical Agricultural Resources, Sanya, 572025, Hainan, China Corresponding author: liping.liu@hitar.org International Journal of Aquaculture, 2024, Vol.14, No.4 doi: 10.5376/ija.2024.14.0018 Received: 15 May, 2024 Accepted: 19 Jun., 2024 Published: 04 Jul., 2024 Copyright © 2024 Wang et al., This is an open access article published under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Preferred citation for this article: Wang Q., Sun C.M., and Liu L.P., 2024, Ecological impacts of common carp invasions: a global perspective, International Journal of Aquaculture, 14(4): 174-183 (doi: 10.5376/ija.2024.14.0018) Abstract As an invasive species that is widely spread globally, common carp (Cyprinus carpio) has caused significant ecological impacts on aquatic ecosystems. The invasion of carp has not only disrupted the ecological balance in many areas, but has also had an economic impact on commercial and recreational fishing, increasing the cost of management and control. This study examines the global status of carp invasion and explores its ecological consequences on the destruction of native species, changes in food web dynamics, and impacts on water quality and sediments. This paper analyzes several cases to show the diversity impacts of carp invasions and the current management strategies, including physical removal, biological control and policy supervision, in order to provide an important theoretical basis for the integrated management of carp invasions and put forward the direction of future research. Keywords Carp invasion; Ecological impacts; Aquatic ecosystems; Biodiversity conservation; Management strategy 1 Introduction The common carp (Cyprinus carpio), originally native to Eurasia, is one of the most widely distributed freshwater fish species globally. Its introduction to various regions has been driven by aquaculture, recreational fishing, and ornamental purposes. The species exhibits traits such as high fecundity, adaptability to diverse environmental conditions, and a generalist diet, which contribute to its success as an invasive species (Crichigno et al., 2016). In regions like Australia, the common carp has become the most abundant large freshwater fish, significantly altering local ecosystems (Koeh, 2004). Similarly, in the Mediterranean region, historical, economic, and cultural factors have facilitated its spread, impacting native aquatic systems (Vilizzi, 2012). Invasive species, such as the common carp, pose significant threats to biodiversity, ecosystem services, and local economies. The global spread of common carp has been well-documented, with established populations in North and South America, Australia, and various parts of Europe and Asia (Nedoluzhko et al., 2021). Ecological niche modeling has shown that common carp can potentially invade a wide range of habitats, from temperate regions to high mountain tropical aquatic systems (Oh et al., 2023). In South America, for instance, the species has expanded its range westward into the Andean region, demonstrating its ability to colonize new environments (Ruppert et al., 2017). The rapid spread and establishment of common carp in diverse regions underscore the need for effective management strategies to mitigate its ecological impacts. This study provides a comprehensive overview of the ecological impacts of carp invasion from a global perspective. To summarize the current distribution and distribution of carp in various regions of the world; assess the ecological consequences of carp invasions on native species and ecosystems; Identify knowledge gaps and propose future research directions to better understand and manage the impacts of common carp invasions. Synthesizing information from multiple studies will contribute to a better understanding of the global ecological impacts of carp and inform policy and management decisions to protect native biodiversity and ecosystem health. 2 Historical Context of Common Carp Invasions 2.1 Origin and natural habitat The common carp (Cyprinus carpio) is native to the Ponto-Caspian and Far Eastern regions, as well as northern
International Journal of Aquaculture, 2024, Vol.14, No.4, 174-183 http://www.aquapublisher.com/index.php/ija 175 Vietnam (Nedoluzhko et al., 2021). Historically, wild populations of genetically pure ancestors are still found in confined areas of Thrace and Northern Anatolia, and possibly in eastern parts of Greece (Vilizzi, 2012). These regions represent the natural habitat where the species evolved and adapted to local environmental conditions. 2.2 Introduction to non-native environments The introduction of common carp to non-native environments has been driven by a combination of historical, economic, and cultural motives. Since Roman times, common carp have been translocated throughout the Mediterranean region for aquaculture and recreational purposes (Vilizzi, 2012). In North and South America, introductions stem from aquaculture facilities and historical introductions for recreational angling. Similarly, in Australia, the species was introduced and has since become the most abundant large freshwater fish in south-east Australia (Crichigno et al., 2016). 2.3 Patterns of spread across different continents The spread of common carp across different continents has followed distinct patterns influenced by environmental conditions and human activities. In the Americas, the distributional potential of common carp covers most temperate regions and high mountain tropical aquatic systems. In southern South America, the species has expanded westward into the Andean Region and established well in the southernmost populations (Vilizzi et al., 2015). In Australia, common carp have spread rapidly and now dominate fish communities over more than 1 million km². The species' success as an invader is attributed to its multiple traits that favor invasion and the degradation of aquatic environments that give it a relative advantage over native species (Samsing et al., 2021). The global perspective on common carp invasions highlights the species' adaptability and the profound ecological impacts it can have on native aquatic faunas. Understanding the historical context and patterns of spread is crucial for developing effective management strategies to mitigate the negative consequences of these invasions. 3 Ecological Impacts on Aquatic Ecosystems 3.1 Disruption of native species 3.1.1 Competition for resources Common carp (Cyprinus carpio) are known to compete with native fish species for resources, significantly impacting their growth and survival. In mesocosm experiments, the presence of common carp reduced the growth of native fish species, although this effect was mitigated by higher native species diversity (Gallardo et al., 2016). Additionally, in Australian dryland rivers, carp monopolized food resources, leading to a decrease in native fish biomass (Marshall et al., 2019). This competition for resources can lead to significant declines in native fish populations, particularly in ecosystems where carp densities are high (Collins et al., 2017). 3.1.2 Predation on native fish Common carp can also directly impact native fish populations through predation. In some ecosystems, carp have been observed to prey on native fish eggs and juveniles, further exacerbating their decline. For example, in Australian dryland rivers, the predation by carp has been linked to the extirpation of an endangered river snail (Marshall et al., 2019). This predatory behavior, combined with their competitive nature, makes common carp a significant threat to native fish species. 3.1.3 Habitat modification Common carp are notorious for their ability to modify habitats, often leading to detrimental effects on native species. Their benthic foraging behavior increases water turbidity and uproots aquatic macrophytes, transforming clear-water lakes into turbid ones (Rolls et al., 2107). This habitat modification not only reduces the availability of suitable habitats for native species but also alters the entire ecosystem's structure and function. In shallow lakes, the presence of common carp has been associated with a shift from macrophyte-dominated clear water states to phytoplankton-dominated turbid states (Matsuzaki et al., 2008). 3.2 Alteration of food web dynamics The introduction of common carp into aquatic ecosystems can lead to significant alterations in food web dynamics.
International Journal of Aquaculture, 2024, Vol.14, No.4, 174-183 http://www.aquapublisher.com/index.php/ija 176 Carp's feeding habits can reduce the abundance of zooplankton and benthic invertebrates, which are crucial components of the aquatic food web (Rolls et al., 2107). This reduction in prey availability can have cascading effects throughout the food web, impacting higher trophic levels, including native fish species. Additionally, the presence of carp can increase phytoplankton abundance due to reduced grazing pressure from zooplankton, further altering the food web structure (Carey and Wahl, 2010). 3.3 Impact on water quality and sediment Common carp significantly impact water quality and sediment composition in invaded ecosystems. Their foraging behavior increases water turbidity and nutrient concentrations, leading to eutrophication and degraded water quality (Marshall et al., 2019). In shallow lakes, carp have been shown to increase suspended solids, phytoplankton, and nutrient levels, contributing to a shift from clear to turbid water states (Carey and Wahl, 2010). These changes in water quality can have far-reaching consequences for the entire aquatic ecosystem, affecting both biotic and abiotic components. 4 Economic and Social Consequences 4.1 Impact on commercial and recreational fishing The invasion of common carp (Cyprinus carpio) has significant repercussions on both commercial and recreational fishing sectors. In Germany, for instance, specialized carp anglers' catch rates exceed commercial harvests by up to 2500%, indicating a substantial impact on the fishing industry. This overabundance of carp can lead to decreased populations of native fish species, thereby reducing the overall biodiversity and altering the ecosystem balance, which in turn affects commercial fishing yields (Busst et al., 2017). Additionally, the presence of carp can shift ecosystems from macrophyte-dominated clear water states to turbid water states dominated by phytoplankton, further complicating fishing activities and reducing the quality of fishing experiences (Gallardo et al., 2016). 4.2 Economic costs of management and control Managing and controlling the spread of common carp incurs significant economic costs. Invasive species like the common carp have been responsible for economic losses amounting to at least US$37.08 billion globally, with North America bearing the highest costs (Haubrock et al., 2021). These costs are associated with damage and resource losses, as well as expenses related to management actions and environmental restoration. In Australia, for example, the control of carp populations in dryland rivers involves substantial investments in biocontrol measures, which are necessary to mitigate their adverse effects on native fish biomass and ecosystem health (Marshall et al., 2019). Moreover, the implementation of continuous removal efforts and novel control approaches is essential to prevent the unintended consequences of increased carp condition and reproductive potential (Coulter et al., 2018). 4.3 Social perceptions and cultural significance The social perceptions and cultural significance of common carp vary across different regions. In some areas, carp are valued for recreational fishing, contributing to local economies through tourism and related activities (Veer and Nentwig, 2015). However, the negative ecological impacts of carp invasions, such as increased water turbidity and reduced native fish populations, can lead to negative social perceptions and a demand for effective management strategies (Figure 1). Policymakers must consider these social welfare implications when deciding on conservation policies and mitigation efforts (Brockmann et al., 2021). Additionally, the cultural significance of carp in certain communities may influence the acceptance and success of management programs, highlighting the need for culturally sensitive approaches to invasive species control. 5 Case Studies of Common Carp Invasions 5.1 North America In North America, the introduction of common carp (Cyprinus carpio) has led to significant ecological changes in various aquatic ecosystems. The carp's presence has been linked to a decline in aquatic plant richness and cover, particularly in the Great Plains and Eastern Temperate Forests ecoregions. Studies have shown that as carp biomass increases, submersed plant cover and species richness decline exponentially, with plant cover reduced to
International Journal of Aquaculture, 2024, Vol.14, No.4, 174-183 http://www.aquapublisher.com/index.php/ija 177 less than 10% and species richness halved in lakes where carp biomass exceeds 190 kg/ha (Bajer et al., 2016). Additionally, the presence of carp has been associated with increased water turbidity and nutrient levels, which further degrade water quality and aquatic habitats. Figure 1 Schematic representation of the Bioeconomic Model. Economic utility or wellbeing is a nested function of space and species-specific decisions, which depend on the linked ecological models-the complex Atlantis model, or the simplified model (Adopted from Brockmann et al., 2021) The impact of common carp on native fish populations has also been documented. For instance, in the Murray-Darling Basin, Australia, carp have been shown to monopolize food resources, leading to a significant reduction in native fish biomass (Marshall et al., 2019). In North America, similar patterns have been observed, with carp outcompeting native species for resources and altering the structure of aquatic communities (Thresher et al., 2018). 5.2Europe In Europe, the introduction of common carp has similarly resulted in ecological disruptions. The carp's benthivorous feeding habits disturb sediment and uproot aquatic plants, leading to increased water turbidity and a decline in macrophyte abundance. This has been observed in various European water bodies, where the presence
International Journal of Aquaculture, 2024, Vol.14, No.4, 174-183 http://www.aquapublisher.com/index.php/ija 178 of carp has led to a shift from clear water states dominated by macrophytes to turbid water states dominated by phytoplankton (Gallardo et al., 2016). The ecological impacts of common carp in Europe are not limited to changes in water quality and plant communities. The introduction of carp has also been associated with the spread of invasive parasites, such as the tapeworm Atractolytocestus huronensis, which has been introduced to Europe along with its host. This parasite poses a threat to both feral and cultured fish populations, highlighting the broader ecological consequences of carp invasions (Costa et al., 2021). 5.3 Australia In Australia, common carp have become a major pest, particularly in the Murray-Darling Basin, where they now constitute up to 90% of fish biomass. The ecological impacts of carp in this region include increased water turbidity, reduced macrophyte density, and changes in macroinvertebrate and native fish assemblages. However, some expected impacts, such as increased turbidity and reduced macroinvertebrate density, were not observed in dryland rivers, suggesting that the general understanding of carp impact requires modification for these specific ecosystems (Figure 2) (Marshall et al., 2019). Efforts to control carp populations in Australia have included the proposed use of Cyprinid Herpesvirus-3 (CyHV-3) as a biological control agent. However, studies have shown that the virus has not yet been detected in the Murray-Darling Basin, and there is little evidence of virus transmission between invasive and native fish species. This highlights the need for careful consideration of ecological context and potential unintended consequences when implementing biocontrol measures (Costa et al., 2021). Figure 2 Network diagram displaying vertebrate-associated viruses identified in native and invasive freshwater fish (Adopted from Costa et al., 2021) Image caption: Colours of each node represent a virus family. Both goldfish and flat-headed gudgeon contained non-vertebrate-associated viruses (Adopted from Costa et al., 2021)
International Journal of Aquaculture, 2024, Vol.14, No.4, 174-183 http://www.aquapublisher.com/index.php/ija 179 6 Management and Control Strategies 6.1 Physical removal and habitat modification Physical removal and habitat modification are common strategies employed to manage invasive common carp populations. Techniques such as netting, electrofishing, and trapping are frequently used to physically remove carp from aquatic ecosystems. For instance, targeting winter aggregations of adult carp for removal has been identified as an effective strategy (Bajer et al., 2019). Habitat modification, such as altering water levels or modifying substrates, can also reduce carp populations by disrupting their breeding and feeding habitats (Coulter et al., 2018). However, these methods often require continuous effort and can be labor-intensive. 6.2 Biological control methods 6.2.1 Use of predators and competitors Biological control through the introduction or enhancement of natural predators and competitors is another strategy. Native predators, such as certain fish species, can help control carp populations by preying on their eggs and larvae (Busst and Britton, 2017). For example, stocking native predators has been successful in controlling juvenile invasive species like the rusty crayfish. However, the effectiveness of this method can vary depending on the specific ecosystem and the presence of suitable predator species. 6.2.2 Sterilization and genetic approaches Sterilization and genetic approaches are emerging as promising methods for controlling invasive carp populations (Feng, 2024). Techniques such as the release of sterilized males or the use of genetic modifications to reduce reproductive success are being explored. In Australia, there are ongoing investigations into the use of a virus to control carp populations. These methods aim to reduce the reproductive capacity of carp, thereby gradually decreasing their numbers over time. 6.2.3 Environmental manipulation Environmental manipulation involves altering the conditions of the habitat to make it less favorable for carp. This can include changes in water quality, such as increasing turbidity or altering nutrient levels, which can negatively impact carp while benefiting native species (Cupp et al., 2021). For example, increasing water turbidity has been shown to reduce carp abundance by disrupting their feeding and breeding behaviors. However, these methods must be carefully managed to avoid unintended consequences on the broader ecosystem. 6.3 Policy and regulatory frameworks Effective management of invasive carp also requires robust policy and regulatory frameworks. These frameworks can include regulations on the transport and release of carp, as well as policies that support the implementation of control measures. Integrated pest management (IPM) plans, which combine multiple control strategies, are being developed by natural resource agencies to address the complex challenges posed by invasive carp (Figure 3) (Kulhanek et al., 2011). Additionally, international cooperation and consistent enforcement of regulations are crucial for preventing the spread of carp across borders and ensuring the success of management efforts (Rytwinski et al., 2019). 7 Ecological and Environmental Considerations 7.1 Long-term ecological effects The long-term ecological effects of common carp (Cyprinus carpio) invasions are profound and multifaceted. Invasive species like common carp can cause significant shifts in ecosystem structure and function. For instance, common carp have been shown to increase water turbidity and nutrient levels, leading to eutrophication and a decline in water quality (Gallardo et al., 2016). These changes can result in stable-state shifts from clear water, macrophyte-dominated systems to turbid, phytoplankton-dominated systems, which are less hospitable to native species. Additionally, the presence of common carp can lead to a reduction in the abundance of benthic macroinvertebrates and submerged macrophytes, further altering the ecosystem. Over time, these changes can lead to a decrease in native fish biomass and biodiversity, as seen in Australian dryland rivers where carp monopolize food resources (Marshall et al., 2019).
International Journal of Aquaculture, 2024, Vol.14, No.4, 174-183 http://www.aquapublisher.com/index.php/ija 180 Figure 3 Underwater acoustic deterrence system (uADS) to assess its effectiveness in reducing the upstream passage of invasive carp (Adopted from Cupp et al., 2021) 7.2 Impact on biodiversity conservation The impact of common carp on biodiversity conservation is significant. Invasive carp can outcompete native species for resources, leading to declines in native fish populations and overall biodiversity (Britton, 2022). For example, in mesocosm experiments, the presence of common carp reduced the growth of native fish species, although this effect was mitigated by higher native species richness. Furthermore, common carp invasions can lead to the extirpation of sensitive species, such as the endangered river snail in Australian rivers, due to predation and habitat alteration (Marshall et al., 2019). The introduction of common carp can also cause genetic introgression and the transmission of non-native pathogens, further threatening native biodiversity. These impacts underscore the importance of managing common carp populations to preserve native biodiversity and ecosystem function. 7.3 Climate change and invasive potential Climate change is expected to exacerbate the invasive potential of common carp. As global temperatures rise, the range of suitable habitats for common carp is likely to expand, increasing the risk of invasion in new areas (Závorka et al., 2018). Climate change can also alter hydrological cycles and nutrient dynamics, creating conditions that favor the establishment and spread of invasive species like common carp. For instance, increased water temperatures and nutrient availability can enhance the growth and reproductive success of common carp, further facilitating their invasion. Additionally, the combined effects of climate change and biological invasions can lead to more severe ecological impacts, such as altered food web dynamics and reduced ecosystem resilience (Shackleton et al., 2018). Therefore, understanding the interactive effects of climate change and invasive species is crucial for predicting and managing future invasions. 8 Concluding Remarks The invasion of common carp (Cyprinus carpio) has been shown to significantly impact aquatic ecosystems globally. Key findings from the reviewed literature indicate that common carp invasions lead to increased water
International Journal of Aquaculture, 2024, Vol.14, No.4, 174-183 http://www.aquapublisher.com/index.php/ija 181 turbidity, elevated nutrient levels, and reduced abundance of macrophytes, zooplankton, and native fish species. These changes often result in a shift from clear water states dominated by macrophytes to turbid water states dominated by phytoplankton, which can have cascading effects on the entire aquatic food web. Additionally, carp invasions have been linked to the decline in species richness and cover of aquatic plants, particularly in regions where carp biomass exceeds critical thresholds. The impacts of carp are not uniform across all ecosystems, with some variations observed based on local environmental conditions and the presence of other stressors. Integrated management approaches are crucial for mitigating the ecological impacts of common carp invasions. Effective management strategies should combine multiple control methods, including biocontrol agents like cyprinid herpesvirus 3 (CyHV-3), physical removal, and habitat modification. The use of integrated pest management (IPM) plans that incorporate behavioral deterrents, barriers, and efficient removal techniques can help limit the spread and reduce the population of invasive carp. Additionally, understanding the specific characteristics of the invaded ecosystems and tailoring management practices to local conditions can enhance the effectiveness of these strategies. Continuous monitoring and adaptive management are essential to address the dynamic nature of carp invasions and to mitigate unintended consequences, such as the potential release from density-dependent competition. Future research should focus on several key areas to improve the management and understanding of common carp invasions. Experts need to conduct more comprehensive studies to examine the long-term ecological impacts of carp in different habitats and environmental conditions. Research should explore the development and implementation of novel biocontrol agents and integrated management technologies, and assess their efficacy and potential ecological consequences; Investigate interactions between carp and other invasive species to understand compounding impacts on native ecosystems. Finally, prediction models that incorporate invasion history, biomass, and environmental variables should be improved to predict the impact of carp invasions and guide management decisions. By filling these research gaps, we can develop more effective strategies to mitigate the ecological impacts of carp and protect aquatic biodiversity. Acknowledgments We sincerely appreciate the opinions of the two peer reviewers, which have made the article more comprehensive. Conflict of Interest Disclosure The authors affirm that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest. Reference Bajer P.G., Beck M.W., Cross T.K., Koch J.D., Bartodziej W., and Sorensen P.W., 2016, Biological invasion by a benthivorous fish reduced the cover and species richness of aquatic plants in most lakes of a large north American ecoregion, Global Change Biology, 22(12): 3937-3947. https://doi.org/10.1111/gcb.13377 Bajer P., Ghosal R., Maselko M., Smanski M., Lechelt J., Hansen G., and Kornis M., 2019, Biological control of invasive fish and aquatic invertebrates: a brief review with case studies, Management of Biological Invasions, 10(2): 227-254. https://doi.org/10.3391/MBI.2019.10.2.02 Britton J., 2022, Contemporary perspectives on the ecological impacts of invasive freshwater fishes, Journal of Fish Biology, 103(4): 752-764. https://doi.org/10.1111/jfb.15240 Brockmann S., Zhang H., Mason D.M., and Rutherford E.S., 2021, Space and species interactions in welfare estimates for invasive species policy, Frontiers in Ecology and Evolution, 9: 703935. https://doi.org/10.3389/fevo.2021.703935 Busst G.M.A., and Britton J.R., 2017, Comparative trophic impacts of two globally invasive cyprinid fishes reveal species‐specific invasion consequences for a threatened native fish, Freshwater Biology, 62(9): 1587-1595. https://doi.org/10.1111/FWB.12970 Carey M., and Wahl D., 2010, Native fish diversity alters the effects of an invasive species on food webs, Ecology, 91(10): 2965-2974. https://doi.org/10.1890/09-1213.1 Collins S.F., Nelson K.A., DeBoom C.S., and Wahl D.H., 2017, The facilitation of the native bluegill sunfish by the invasive bighead carp, Freshwater Biology, 62(9): 1645-1654. https://doi.org/10.1111/FWB.12976
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International Journal of Aquaculture, 2024, Vol.14, No.4, 184-194 http://www.aquapublisher.com/index.php/ija 184 Review Article Open Access Molecular Mechanisms Underlying the Diversification of Aquatic Life Forms ManmanLi Hainan Institute of Biotechnology, Haikou, 570206, Hainan, China Corresponding email: 502684238@qq.com International Journal of Aquaculture, 2024, Vol.14, No.4 doi: 10.5376/ija.2024.14.0019 Received: 20 May, 2024 Accepted: 30 Jun., 2024 Published: 15 Jul., 2024 Copyright © 2024 Li., This is an open access article published under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Preferred citation for this article: Li M.M., 2024, Molecular mechanisms underlying the diversification of aquatic life forms, International Journal of Aquaculture, 14(4): 184-194 (doi: 10.5376/ija.2024.14.0019) Abstract The diversification of aquatic life forms is a significant phenomenon in biological evolution, and understanding the molecular mechanisms behind it is crucial for uncovering the origins of biodiversity and its adaptability. This study explores the key molecular mechanisms driving the diversification of aquatic life forms, including gene duplication, horizontal gene transfer, mutations, and their roles in genetic variability. It also analyzes how gene regulatory networks and epigenetic modifications influence the evolution of developmental pathways, and how environmental factors such as water conditions, predation pressure, and climate change drive molecular diversification. By examining the molecular evolution of key aquatic adaptations such as respiratory systems, sensory systems, and reproductive strategies, the study summarizes the evolutionary strategies of aquatic organisms in various ecological environments. These findings not only deepen the understanding of biodiversity and adaptive evolution in aquatic organisms but also provide a scientific basis for the development of biodiversity conservation strategies and highlight future research directions. Keywords Diversification; Molecular mechanisms; Aquatic life forms; Gene duplication; Evolutionary strategies 1 Introduction Aquatic life forms encompass a vast array of organisms that inhabit marine, freshwater, and brackish environments. These organisms have evolved a multitude of adaptations to thrive in their respective habitats, ranging from physiological mechanisms to morphological traits. For instance, water striders have developed superhydrophobic bristles and elongated legs to navigate water surfaces, a transition that occurred approximately 200 million years ago. Similarly, the Nepomorpha, or true water bugs, have diversified significantly since the late Permian, adapting to various aquatic environments. The deep-sea fish Coryphaenoides rupestris exhibits genotypic segregation by depth, highlighting the role of vertical habitat gradients in promoting intraspecific diversity (Gaither et al., 2018; Ye et al., 2020; Finet et al., 2022). Understanding the mechanisms underlying the diversification of aquatic life forms is crucial for several reasons. It provides insights into the evolutionary processes that drive speciation and adaptation. For example, the study of carbon concentrating mechanisms (CCMs) in aquatic photosynthetic organisms reveals how these systems contribute to global net primary productivity and marine carbon sequestration (Griffiths et al., 2017; Zhou and Mai, 2024). It helps in predicting how current and future environmental changes, such as global warming and cooling, will impact biodiversity. Research on the Anomura crustaceans shows that speciation rates are influenced by global temperature changes, with marine clades responding differently to cooling and warming trends (Davis et al., 2016). Understanding diversification aids in conservation efforts by identifying key adaptive traits and environmental pressures that shape biodiversity. This study leverages current knowledge on the molecular mechanisms driving the diversification of aquatic life forms to analyze the genetic and molecular basis of key adaptive traits in various aquatic organisms. It explores the role of environmental factors, such as climate change and habitat alterations, in shaping diversification patterns and discusses the implications of these findings for biodiversity conservation and management.By
International Journal of Aquaculture, 2024, Vol.14, No.4, 184-194 http://www.aquapublisher.com/index.php/ija 185 integrating the results of multiple studies, this research aims to provide a comprehensive understanding of how molecular mechanisms promote the diversification of aquatic life forms, thereby offering valuable insights for evolutionary biology and conservation science.. 2 Genetic Basis of Diversification 2.1 Role of gene duplication Gene duplication is a fundamental mechanism that contributes to the diversification of aquatic life forms by providing raw genetic material for evolutionary innovation. For instance, the genome of the water strider Gerris buenoi reveals numerous local gene duplications and expansions of gene families that are crucial for adaptations to a water surface lifestyle. These duplications affect processes such as growth, vision, desiccation resistance, detoxification, olfaction, and epigenetic regulation, which are essential for the water strider's survival and diversification in aquatic environments (Armisén et al., 2018). Similarly, the duplication of the Na+/K+-ATPase gene in clitellate annelids is associated with their transition from marine to freshwater habitats, suggesting that these duplications played a significant role in their diversification (Horn et al., 2019). The dynamic evolution of gene families, including gene duplication and deletion, has been shown to drive phenotypic diversity and adaptation in various aquatic species, such as African cichlid fishes and Hawaiian drosophilids (Harris and Hofmann, 2015). 2.2 Horizontal gene transfer Horizontal gene transfer (HGT) is another critical mechanism that accelerates microbial evolution and promotes diversification and adaptation in aquatic environments. In marine ecosystems, the cyanobacterium Prochlorococcus exhibits a highly streamlined genome with frequent gene exchange, facilitated by mobile genetic elements known as tycheposons. These elements drive genomic plasticity and niche differentiation by carrying genes important for nutrient acquisition and other adaptive traits. The efficient dispersal and transmission of tycheposons through extracellular vesicles and phage particles highlight the significant role of HGT in the diversification of marine microbes (Hackl et al., 2020). Furthermore, the formation of new ecologically distinct populations (ecotypes) in the Microcystis aeruginosa complex is driven by mutations in key functional genes and the acquisition of new metabolic pathways through HGT, allowing these populations to exploit new resources and coexist with parental populations (Escalera et al., 2021). 2.3 Mutations and genetic variability Mutations and genetic variability are essential for the adaptive diversification of aquatic organisms. In the rotifer Brachionus calyciflorus, genetic variation and differences in gene expression related to temperature tolerance play a significant role in local adaptation and ecological diversification. The identification of candidate genes associated with metabolism and stress response highlights the importance of genetic variability in adapting to different thermal environments (Paraskevopoulou et al., 2019). The adaptation of fishes to changing salinity involves genetic changes in osmoregulatory systems, with natural selection targeting genes coding for key cellular ion exchange enzymes such as V-type, Ca2+, and Na+/K+-ATPases. These genetic changes support the transition across salinity boundaries and contribute to the diversification of fish species (Velotta et al., 2022). The study of diatoms also reveals that habitat transitions, such as from marine to freshwater environments, are influenced by genetic variability and adaptive mutations, which play a crucial role in their diversification (Roberts et al., 2023). 3 Regulatory Networks and Evolution 3.1 Gene regulatory networks Gene regulatory networks (GRNs) play a crucial role in the diversification of aquatic life forms by orchestrating the expression of genes in response to environmental changes. In marine mammals, for instance, the adaptation to aquatic environments involved significant modifications in GRNs. Comparative genomics has revealed that genes associated with thermoregulation, such as NFIA and SEMA3E, have undergone convergent evolution to facilitate the transition from land to water. These genes are involved in the formation of blubber and vascular development, respectively, which are essential for maintaining body temperature in aquatic environments (Figure 1) (Triant et
International Journal of Aquaculture, 2024, Vol.14, No.4, 184-194 http://www.aquapublisher.com/index.php/ija 186 al., 2021; Yuan et al., 2021). The loss of the UCP1 locus in many marine mammals indicates a shift in their thermostatic strategy from enhancing heat production to limiting heat loss, further highlighting the role of GRNs in evolutionary adaptation. The gene plays a crucial role in determining whether adipocytes develop into energy-storing white fat cells or heat-producing brown fat cells by regulating adipocyte differentiation. In brown fat cells, the action of the UCP1 protein can convert stored energy into heat, which is vital for maintaining body temperature in cold aquatic environments. Research has found convergent amino acid substitutions in the NFIA gene across multiple marine mammal species, suggesting that these species may promote the formation of adipose tissue through similar mechanisms to adapt to aquatic environments. Additionally, the SEMA3Egene plays an important role in vascular development. The vascular structures of marine mammals, such as the complex retia mirabilia, are crucial for thermoregulation, effectively reducing heat loss in water. Studies show that the convergent evolution of the SEMA3E gene may help these animals develop their vascular systems more effectively to maintain body temperature in cold environments. Figure 1 Convergent evolution of thermoregulation in marine mammals (Adopted from Yuan et al., 2021) Image caption: (A) Schematic diagram of thermoregulation in marine mammals. Up- or down-regulation of nuclear factor I A (NFIA) affects the cell fate of mesenchymal precursors, the integrity of UCP1 gene affects the fate of brown adipocyte, and the well-developed retia mirabilia in marine mammal aids in the heat transfer to maintain body temperature balance. VSMC, vascular smooth muscle cell; EC, endothelial cell. (B) A unique amino acid change in the NFIAgene of marine mammals. Shared amino acid change are highlighted in blue, IP, Indo-Pacific. (C) A unique amino acid change in the SEMA3E gene of cetaceans and pinnipeds. Blue highlighting indicates the shared amino acid change. (D) VISTA sequence conservation plot of the UCP1 gene, using goat (ARS1) as a reference (Adopted from Yuan et al., 2021) 3.2 Epigenetic modifications Epigenetic modifications, including DNA methylation and histone modifications, are pivotal in regulating gene expression without altering the underlying DNA sequence. These modifications can lead to phenotypic changes that are essential for the adaptation to new environments. In the context of aquatic life forms, epigenetic changes may influence the expression of genes involved in critical physiological processes such as thermoregulation, diving, and navigation. For example, the unique changes in genes like NFIA and SEMA3E in marine mammals
International Journal of Aquaculture, 2024, Vol.14, No.4, 184-194 http://www.aquapublisher.com/index.php/ija 187 suggest that epigenetic mechanisms could be at play in regulating these genes to meet the demands of an aquatic lifestyle (Yuan et al., 2021). Although specific studies on epigenetic modifications in aquatic life forms are limited, the existing evidence underscores their potential role in facilitating rapid and reversible adaptations to environmental changes. 3.3 Evolution of developmental pathways The evolution of developmental pathways is a key factor in the diversification of aquatic life forms. Developmental pathways are regulated by a complex interplay of transcription factors and gene regulatory networks. In plants, for example, the evolution from unicellular aquatic algae to complex flowering plants involved significant changes in transcription factors and their associated regulatory networks. These changes were driven by alterations in DNA binding specificity, protein-protein interactions, and cis-regulatory elements, which collectively contributed to the functional evolution of transcription factors (Romani and Moreno, 2020). Similarly, in marine mammals, the adaptation to aquatic environments required modifications in developmental pathways to support new physiological and morphological traits. The study of these evolutionary processes in both plants and animals provides valuable insights into the molecular mechanisms underlying the diversification of life forms in aquatic environments. 4 Environmental Factors Driving Molecular Diversification 4.1 Adaptation to different water conditions Adaptation to varying water conditions is a significant driver of molecular diversification in aquatic life forms. The transition between marine and freshwater environments, characterized by steep salinity gradients, poses a substantial osmotic challenge for many species. For instance, diatoms exhibit molecular acclimation mechanisms to cope with hypo-osmotic stress when transitioning from saline to freshwater environments, highlighting the role of cellular mechanisms in overcoming salinity barriers (Bilcke and Kamakura, 2023). Similarly, the genomic adaptations in marine mammals, such as changes in genes associated with thermoregulation and deep diving, underscore the importance of molecular changes in adapting to aquatic lifestyles. These adaptations are crucial for survival and diversification in different aquatic habitats. 4.2 Pressure of predation and competition Predation and competition are pivotal biotic factors influencing the diversification of aquatic species. In postglacial freshwater fish species, predation and competition have been implicated in the divergence into multiple ecomorphs within lakes, suggesting that these biotic pressures drive morphological and ecological diversification (Condamine et al., 2019; Tiddy et al., 2023). The unequal body shape diversification in the Gondwanan fish radiation (Characiformes) is influenced by competition for niches and habitat variation. Neotropical characiform lineages exhibit greater morphological diversity compared to their African counterparts, likely due to competition with cypriniform fishes in Africa and the availability of diverse habitats in the Neotropics (Burns et al., 2023). These examples illustrate how predation and competition pressures can lead to significant diversification in aquatic life forms. 4.3 Influence of climate change Climate change is a critical abiotic factor driving molecular diversification in aquatic ecosystems. Global cooling and warming have been shown to influence speciation rates in marine and freshwater clades differently. For example, speciation rates in marine clades of Anomura crustaceans are positively correlated with global cooling, while freshwater clades show increased speciation rates with global warming (Davis et al., 2016). Climate fluctuations have played a significant role in the diversification of true water bugs (Nepomorpha) during the Mesozoic, with palaeoecological opportunities promoting lineage diversification (Friedman et al., 2021). The interaction of multiple climate-change drivers, such as ocean warming, acidification, and deoxygenation, also affects aquatic productivity and species responses, highlighting the complex influence of climate change on molecular diversification (Häder and Gao, 2023). Understanding these dynamics is crucial for predicting and mitigating the impacts of ongoing climate change on aquatic biodiversity.
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